Myrmecologische Nachrichten 8 271 - 276 Wien, September 2006

Change in the zoogeographical structure of (: Formicidae) caused by urban pressure in the Sofia region (Bulgaria)

Vera ANTONOVA & Lyubomir PENEV

Abstract A comparison of the zoogeographical peculiarities of the myrmecofauna in urban and non-urban habitats in Sofia re- gion is performed. The proportion of the widespread Holarctic species was twice as high in the urban as compared to the non-urban sites. Also Euro-Caucasian elements are well represented in the urban sites. The share of South-Transpalae- arctic, Eurasian steppe and Boreo-Montane elements is low or such elements are absent from the urban sites. Key words: Ants, Formicidae, urban landscape, Bulgaria, zoogeography. Vera Antonova (contact author) & Dr. Lyubomir Penev, Central Laboratory of General Ecology, Bulgarian Academy of Sciences, 2 Gagarin Str., 1113 Sofia, Bulgaria. E-mail: [email protected]

Introduction Comparatively few investigations exist on the change of The main anthropogenic factors are: disturbance of the struc- zoogeographical structure of ants in urban areas. It is well ture of habitats, changes in microclimate and nutrition re- known that widespread invertebrate species are common in sources (ANDERSEN 1990, 1995a). Unfortunately, most of urban environments compared to its surroundings (KLAUS- the species of ants from the above mentioned studies do not NITZER 1990). This pattern has been observed for ants in exist in the European myrmecofauna and thus direct com- different parts of the world: Japan – Tokyo and Chiba (KON- parisons are not possible. DOH 1978, YAMAGUCHI 2004), Kitakyushu (KONDOH & KI- In this study we investigate changes in the zoogeo- TAZAWA 1984); USA – Raleigh (North Carolina) (NUHN & graphical structure of the fauna in both urban and rural WRIGHT 1979), urban zones on the Atlantic coast (KING landscape of the city of Sofia. We discuss the possibility & GREEN 1995 – in MCINTYRE 2000); Europe – Warsaw of using the zoogeographical structure of ants for bioindi- (PISARSKI & CZECHOWSKI 1978, BANASZAK & al. 1978, cation in urban landscapes. CZECHOWSKI & al. 1979, PISARSKI 1982, PISARSKI & CZE- Material and methods CHOWSKI 1987, CZECHOWSKI & al. 1990, CZECHOWSKI & PISARSKI 1990, CZECHOWSKI 1991), Espoo (VEPSÄLÄIN- The city of Sofia is situated at 550 m a.s.l. on average, the EN & WUORENRINNE 1978), Liege (GASPAR & THIRION average annual temperature is around 10.2 °C and the 1978), Garraf (Barcelona) (RESTREPO & al. 1985), Leipzig average annual rainfall is around 560 mm (BLUSKOVA & 2 (RICHTER & al. 1986), Köln (LIPPKE & CÖLLN 1991, BEHR al. 1983). The urban area covers approximately 220 km & al. 1996), Gönnersdorf (BEHR & CÖLLN 1993), Mainz (KOVACHEV 2005). The central part of the city is situated (DAUBER 1995, DAUBER 1997, DAUBER & EISENBEIS 1997), at approximately 42° 41' N and 23° 19' E. With respect to Wien (SCHLICK-STEINER & STEINER 1999), Linz (AMBACH climate, Sofia is situated relatively near (c. 120 km) to the 1999). Generally, each urban fauna is a selection from the border between temperate and submediterranean climates. regional species pool, therefore, depending on a city's geo- The material for the present study was collected in graphical position, species from the respective zoogeogra- 2003 - 2005. It is kept in the collection of V. Antonova in phical zone prevail (PISARSKI & KUĹESZA 1982, VEPSÄLÄ- the Central Laboratory of General Ecology of the Bulgari- INEN & PISARSKI 1982). PISARSKI & VEPSÄLÄINEN (1987) an Academy of Sciences. Fifty-six of the localities were note that the share of Palaearctic ants is up to 90 % in Eur- situated in the urban greenery ("urban" sites in Fig. 1): opean cities of the temperate climatic zone while that of wooded and open areas in parks, green yards along trans- European species is relatively small. In Mainz European port corridors and streets; amounting to a total of ten "for- and Holarctic ant species prevail (DAUBER 1995). In War- est" sites and 46 "meadow" sites (Fig. 1). Another eight lo- saw Palaearctic and South-Eurosiberian ants constitute the calities are situated in the vicinity of Sofia ("non-urban" bulk of the fauna (PISARSKI & CZECHOWSKI 1987). sites in Fig. 1), at the foot of Vitosha, Lyulin and Lozen Many authors studied the possibility to use ants as bio- Mountains; amounting to a total of two "forest" sites and indicators on the basis of functional groups (ANDERSEN six "meadow" sites. The non-urban sites are situated not 1990, 1995a, b, 1997a, b, c, d, ANDERSEN & SPARLING 1997, higher than 150 m above the average altitude of the city VANDERWOUDE & al. 1997а, b, CHUDZICKA & SKIBIŃSKA with distance up to 10 km from the city ring road. 1998, PECK & al. 1998, NEW 2000, READ & ANDERSEN The urban forest localities are situated within the Sofia 2000, HOFFMANN & al. 2000, ANDERSEN & al. 2002, HOFF- GLOBENET sites described by PENEV & al. (2004). Spe- MANN & ANDERSEN 2003). Anthropogenic disturbances cies compositions of plant communities are mostly an- have a different impact on different ant species, dependent thropogenic. Most of the sites are situated in oak forests on their ecological needs (PĘTAL 1980, ANDERSEN 2000). (50 - 90 years old) with prevalence of the mesophilous in-

Stefan-Schödl-Gedenkband / Stefan Schödl Memorial Volume

Fig. 1: Sampling sites in Sofia city and surroundings. M - meadow; F – forest; ■ - non-urban site; ● – urban site. troduced Red Oak (Quercus rubra L.) (formation Quer- SOV & DLUSSKY (1992), SEIFERT (1996), and CZECHOWS- ceta rubrae). The sites in open urban areas were chosen KI & al. (2002). on the basis of prevalence of the dominant plant species: The zoogeographical classification of ants was taken Poa spp., Festuca spp., Phleum pratense L., Avena fatua from ATANASSOV (1952) and CZECHOWSKI & al. (2002), L., Setaria viridis (L.), Cynodon dactylon PERS., Dactylis with some changes and additions. glomerata L., Trifolium spp., Taraxacum officinale WEB- The reliability of differences in percentage within each BER, Bellis perennis L., Capsella bursa-pastoris MED., and zoogeographical category between the city and its vicinity Plantago spp. Similar plant communities were chosen a- was tested with the "Differences between two percentag- round the city, in the non-urban sites (Fig. 1). es" two-sided significance test available in Statistica for For the purpose of the present study the direct sampl- Windows 5.0 (STATSOFT 1995), which is suitable for ana- ing method (BESTELMEYER & al. 2000) was used. In the for- lysing data from uneven sampling design. est sites ant colonies were searched in the leaf litter, in ______the ground (up to 25 cm depth), under stones, under the Tab. 1: Zoogeographical classification of the ant species in bark of trees, in logs and nuts. In the meadow sites colo- Sofia and its surroundings. BM – Boreo-Montane; NP – nies were searched in the ground, in the grass turf and un- North-Transpalaearctic; H – Holarctic; AP – Amphipalae- der stones. The sampling area per site was approximately arctic; E – European; ES – Euro-Siberian s.str.; EC – Euro- 1 ha and search was conducted for 4 hours per site. The to- Caucasian; CE – Central-European; SP – South-Trans- tal number of colonies we found is 1634. Ants were de- palaearctic; St – Eurasian steppe; Pont – Pontic; MD – termined to species level by use of the keys of ATANAS- Mediterranean s.str. →

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No. Species Zoogeographical element Sofia Surroundings

1. Ponera coarctata (LATREILLE, 1802) MD + 2. rubra (LINNAEUS, 1758) H + 3. Myrmica ruginodis NYLANDER, 1846 NP + + 4. Myrmica rugulosa NYLANDER, 1849 ES + + 5. NYLANDER, 1846 H + + 6. Myrmica schencki VIERECK, 1903 H + + 7. Myrmica specioides BONDROIT, 1918 CE + 8. Myrmica sulcinodis NYLANDER, 1846 BM + + 9. Myrmica rugulososcabrinodis KARAWAJEW, 1929 Pont + 10. Myrmica lonae FINZI, 1926 ES + + 11. Myrmica salina RUZSKY, 1905 ES + 12. Myrmica sabuleti MEINERT, 1861 ES + + 13. debile (FOERSTER, 1850) EC + 14. Solenopsis fugax (LATREILLE, 1798) MD + + 15. Temnothorax nylanderi (FOERSTER, 1850) CE + + 16. Temnothorax affinis (MAYR, 1855) EC + 17. Temnothorax tuberum (FABRICIUS, 1775) ES + + 18. Temnothorax unifasciatus (LATREILLE, 1798) EC + + 19. Messor structor (LATREILLE, 1798) MD + 20. cf. caespitum (LINNAEUS, 1758) H + + 21. Tetramorium hungaricum RÖSZLER, 1935 MD + 22. Tetramorium moravicum KRATOCHVIL, 1941 ? + + 23. Myrmecina graminicola (LATREILLE, 1802) AP + + 24. Dolichoderus quadripunctatus (LINNAEUS, 1771) ES + + 25. Tapinoma erraticum (LATREILLE, 1798) MD + + 26. Plagiolepis pygmaea (LATREILLE, 1798) MD + + 27. Plagiolepis taurica SANTSCHI, 1920 St + 28. Camponotus vagus (SCOPOLI, 1763) ES + + 29. Camponotus aethiops (LATREILLE, 1798) MD + + 30. Camponotus ligniperda (LATREILLE, 1802) E + 31. Camponotus piceus (LEACH, 1825) MD + + 32. Camponotus fallax (NYLANDER, 1856) ES + + 33. Camponotus truncates (SPINOLA, 1808) EC + 34. Prenolepis nitens (MAYR, 1853) Pont + 35. niger (LINNAEUS, 1758) H + + 36. Lasius platythorax SEIFERT, 1991 NP? + + 37. Lasius brunneus (LATREILLE, 1798) EC + + 38. Lasius alienus (FOERSTER, 1850) H + + 39. Lasius paralienus SEIFERT, 1992 E + + 40. Lasius psammophilus SEIFERT, 1992 E + 41. Lasius flavus (FABRICIUS, 1782) H + + 42. Lasius citrinus EMERY, 1922 SP + + 43. Lasius balcanicus SEIFERT, 1988 ? + 44. Lasius fuliginosus (LATREILLE, 1798) AP + + 45. Formica fusca LINNAEUS, 1758 NP + + 46. Formica cinerea MAYR, 1853 EC + + 47. Formica rufibarbis FABRICIUS, 1793 ES + + 48. Formica cunicularia LATREILLE, 1798 EC + + 49. Formica gagates LATREILLE, 1798 SP + + 50. Formica pratensis RETZIUS, 1783 SP + + 51. Formica rufa LINNAEUS, 1761 NP + + 52. Formica sanguinea LATREILLE, 1798 SP + + 53. Polyergus rufescens (LATREILLE, 1798) EC + + 54. Cataglyphis nodus (BRULLÉ, 1832) St +

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Figs. 2 - 4: Zoogeographical structure of the ants, based on frequency of occurrence of species, of (2) all sites of the stud- ied region, (3) the sites in the urban area, and (4) the sites in the non-urban area. BM – Boreo-Montane; NP – North- Transpalaearctic; H – Holarctic; AP – Amphipalaearctic; E – European; ES – Euro-Siberian s.str.; EC – Euro-Caucasian; CE – Central-European; SP – South-Transpalaearctic; St – Eurasian steppe; Pont – Pontic; MD – Mediterranean s.str.

Tab. 2: Species with significant differences between their frequencies of occurrence in urban and non-urban areas.

Species Zoogeographical Frequency of oc- Frequency of oc- P-level of element currence in Sofia currence in vicinity differences Camponotus aethiops Mediterranean s.str. 0 37.5 0.0000 Camponotus ligniperda European 0 37.5 0.0000 Camponotus piceus Mediterranean s.str. 1.78 37.5 0.0003 Messor structor Mediterranean s str. 0 25.0 0.0003 Formica rufa North-Transpalaearctic 0 25.0 0.0003 Plagiolepis taurica Eurasian steppe 0 25.0 0.0003 Cataglyphis nodus Eurasian steppe 0 25.0 0.0003 Myrmica schencki Holarctic 5.40 37.5 0.0035 Formica sanguinea South-Transpalaearctic 1.78 25.0 0.0050 Formica gagates South-Transpalaearctic 0 12.5 0.0085 Myrmica sulcinodis Boreo-Montane 0 12.5 0.0098 Lasius citrinus South-Transpalaearctic 19.60 62.5 0.0102 Formica pratensis South-Transpalaearctic 8.90 37.5 0.0242

Results and discussion We counted the frequency of occurrence of each spe- cies for the urban and non-urban sites separately. On that The number of species found in the studied region is 54 basis we summarized the percentage share of the zoogeo- (Tab. 1). Among them 44 (81 %) belong to the Euro-Sibe- graphical categories for the urban and non-urban areas se- rian complex, 8 (15 %) to the Mediterranean complex and parately. The share of the Holarctic species is almost twice the zoogeographical status of 2 species (4 %) was not pro- as high in the urban area (39 %) in comparison to the non- perly identified (Fig. 2). urban area (19 %), which is illustrated by Figures 3 and 4. Within the Euro-Siberian complex, the highest percen- Also Euro-Caucasian elements are more frequent in urban tages (by number of species) are represented by Euro- (24 %) compared to non-urban sites (15 %). Figure 3 shows Siberian s.str. (16 %), Euro-Caucasian (14 %) and Hol- the uneven representation of the particular zoogeographi- arctic (13 %) elements. The other categories of the same cal categories in the urban area while in Figure 4, non-urban complex are represented by less than 4 species (7 %, Fig. 2). area, the proportions are relatively balanced. The most There are only 8 (15 %) Mediterranean s.str. species. Con- widespread Holarctic species in Sofia that are more fre- sidering that the total percentage of Mediterranean ant spe- quent in urban than in non-urban sites are Tetramorium cf. cies in Bulgaria is 34.3% (ATANASSOV & DLUSSKY 1992), caespitum (LINNAEUS, 1758) (77 % frequency of occur- the percentage of this group is relatively low in our study rence in urban and 62 % in non-urban sites) and Lasius area. The small share of Mediterranean species is probably niger (LINNAEUS, 1758) (66 % in urban and 50 % in non- caused by the relatively high altitude of the city of Sofia urban zone). Euro-Caucasian species with higher frequen- situated in a kettle surrounded by mountains. cy of occurrence in the urban than in the non-urban zone

274 are: Formica cinerea MAYR, 1853 (61 % in urban and 50 % cipe. – Proceedings of the Ecological Society of Australia 16: in non-urban zone), Temnothorax affinis (MAYR, 1855) 347-357. (11 % in urban and 0 % in non-urban zone) and Stenamma ANDERSEN, A.N. 1995a: A classification of Australian ant com- debile (FOERSTER, 1850) (5 % in the urban and 0 % in the munities, based on functional groups which parallel plant life- non-urban zone). Hence, the most widespread species re- forms in relation to stress and disturbance. – Journal of Bio- present the bulk of the myrmecofauna in the urban area. geography 22: 15-29. This assertion is in accordance with other similar studies on ANDERSEN, A.N. 1995b: Measuring more of biodiversity: genus urban myrmecofaunas (PISARSKI & KUĹESZA 1982, VEP- richness as a surrogate of species richness in Australian ant SÄLÄINEN & PISARSKI 1982, PISARSKI & CZECHOWSKI faunas. – Biological Conservation 73: 39-43. 1987, KLAUSNITZER 1990). ANDERSEN, A.N. 1997a: Ants as indicators of ecosystem restor- Among all species, only thirteen have statistically sig- ation following mining: a functional group approach. In: HALE, nificant differences in their frequency of occurrence (Tab. P. & LAMB, D. (Eds.): Conservation Outside Nature Reserves. 2). We may conclude that the share of these thirteen spe- – Centre for Conservation Biology, University of Queensland, pp. 319-325. cies decreases in – and some are even absent from – the ur- ban area. All South-Transpalaearctic, Eurasian steppe and ANDERSEN, A.N. 1997b: Functional groups and patterns of or- Boreo-Montane species found in the studied region are ganization in North American ant communities: a compar- ison with Australia. – Journal of Biogeography 24: 433-460. more abundant in the non-urban area. Among the other zoo- geographical categories only single species differ in their ANDERSEN, A.N. 1997c: Using ants as bioindicators: Multiscale frequencies of occurrence (3 Mediterranean, 1 North-Trans- issues in ant community ecology. – Conservation Ecology [on- line] 1: 8. palaearctic, 1 Holarctic and 1 European species). Thus, the species mentioned in Table 2 may be considered as indi- ANDERSEN, A.N. 1997d: Measuring invertebrate biodiversity: sur- cators for the non-urban zones in the studied region. They rogates of ant species richness in the Australian seasonal trop- ics. – Memoirs of the Museum of Victoria 56: 355-359. show a weak tolerance to urban pressure. We may conclude that the prevalence of widespread ANDERSEN, A.N. 2000: Global ecology of rainforest ants. In: Holarctic species in the urban area may serve as indicator AGOSTI, D., MAJER, J.D., ALONSO, L.E. & SCHULTZ, T. (Eds.): Ants: standard methods for measuring and monitoring biodi- of urban pressure. Most of these species can be charac- versity. – Smithsonian Institution Press, Washington and Lon- terized by wide ecological tolerance and this may explain don, pp. 25-34. their prevalence in almost all urban habitats. The small ANDERSEN, A.N., HOFFMANN, B.D., MÜLLER, J.W. & GRIFFITHS, share or the absence of South-Transpalaearctic, Eurasian A.D. 2002: Using ants as bioindicators in land management: steppe and Boreo-Montane elements may be also a conse- simplifying assessment of ant community responses. – Jour- quence for urban pressure. Thus, the zoogeographical struc- nal of Applied Ecology 39: 8-17. ture may be used for purposes of bioindication. Further ANDERSEN, A.N. & SPARLING, G.P. 1997: Ants as indicators of studies in regions with similar climate conditions are need- restoration success: relationship with soil microbial biomass ed to strengthen this evidence. in the Australian seasonal tropics. – Restoration Ecology 5: 109-114. Acknowledgements ATANASSOV, N. 1952: Regularities in the distribution and biolo- This work was partly supported by National Science Fund gical observations on the ants in Vitosha mountain. – Trudove – Bulgarian Ministry of Education and Science (grant No. na planinska prirodonauchna stantzia Vitosha 1: 1-214 (in B-1305/03). We are deeply grateful to Prof. Dr. Wojciech Bulgarian). Czechowski, Dr. Wieslawa Czechowska and Prof. Dr. Rad- ATANASSOV, N. & DLUSSKY, G. 1992: Fauna Bulgarica. 22. Hy- chenko for the salutary taxonomical discussions. We are menoptera, Formicidae. – Bulgarian Academy of Sciences, So- very grateful to the editors for the valuable recommenda- fia, 310 pp. (in Bulgarian). tions and to both referees for the critical remarks. BANASZAK, J., CZECHOWSKI, W., PISARSKI, B., & SKIBINSKA, E. 1978: Owady społeczne w środowisku zurbanizowanym. – Zusammenfassung Kosmos (Warsaw) 27: 173-180. Zoogeographische Aspekte der Ameisenfauna der Region BEHR, D. & CÖLLN, K. 1993: Zur Ameisenfauna (Hymenoptera, Sofia werden behandelt. Insbesondere werden urbane und Formicidae) von Gönnersdorf (Kr. Daun). – Dendrocopos 20: nicht-urbane Lebensräume verglichen. Der Anteil weitver- 148-160. breiteter holarktischer Arten war in den urbanen Lebens- BEHR, D., LIPPKE, S. & CÖLLN, K. 1996: Zur Kenntnis der Amei- räumen doppelt so groß wie in den nicht-urbanen. Desglei- sen von Köln (Hymenoptera, Formicidae). – Decheniana-Bei- chen waren euro-kaukasische Elemente in der urbanen Zone hefte 35: 215-232. gut vertreten. Hingegen ist der Anteil süd-transpaläarkti- BESTELMEYER, B.T., AGOSTI, D., ALONSO, L.E., BRANDÃO, R.F., scher, eurasischer Steppen- und boreo-montaner Elemente BROWN, JR.W.L., DELABIE, H.C.J. & SILVESTRE, R. 2000: Field in der urbanen Zone niedrig, oder diese Elemente fehlen techniques for the study of ground-dwelling ants: An overview, hier völlig. description and evaluation. In: AGOSTI, D., MAJER, J.D., ALON- SO, L.E. & SCHULTZ, T. (Eds.): Ants: standard methods for mea- suring and monitoring biodiversity. – Smithsonian Institution References Press, Washington and London, pp. 122-144. AMBACH, J. 1999: Verbreitung der Ameisenarten (Hymenoptera: BLUSKOVA, D., ZLATKOVA, L., LINGOVA, ST., MODEVA, ZH., SU- Formicidae) im Linzer Stadtgebiet (Oberösterreich) und ihre BEV, L. & TENEVA, M. 1983: Climate and microclimate of So- Bewertung aus stadtökologischer Sicht. – Naturkundliches Jahr- fia. – Bulgarian Academy of Sciences, Sofia, 154 pp. (in Bul- buch der Stadt Linz 44: 191-320. garian). ANDERSEN, A.N. 1990: The use of ant communities to evaluate CHUDZICKA, E. & SKIBIŃSKA, E. 1998: Monitoring and role of change in Australian terrestrial ecosystems: a review and a re- terrestrial invertebrates in bioindicatory evaluation of en-

275 vironment condition and changes. – Memorabilia Zoologica PENEV, L.D., STOYANOV, I.L., DEDOV, I., DIMITROV, D., GROZEVA, 51: 3-12. M. & DOICHINOVA, V. 2004: The Sofia (Bulgaria) GLOBE- CZECHOWSKI, W. 1991: Comparison of the myrmecofaunas (Hy- NET sites: Description spatial variation of the landscape mosa- menoptera, Formicoidea) of tree stands and lawns in Warsaw ic. In: PENEV, L., NIEMELA, J., KOTZE, J. & CHIPEV, N. (Eds.): parks. – Fragmenta Faunistica 35: 179-183. Ecology of the city of Sofia. Species and communities in ur- ban environment. – PENSOFT Publishers, Sofia, Moscow, pp. CZECHOWSKI, W., CZECHOWSKA, W. & PALMOWSKA, J. 1990: Ar- 3-24. boreal myrmecofauna of Warsaw parks. – Fragmenta Faunis- tica 34: 37-45. PĘTAL, J. 1980: Ant populations, their regulation and effect on soil in meadows. – Ekologia Polska 28: 297-326. CZECHOWSKI, W. & PISARSKI, B. 1990: Ants (Hymenoptera, For- micidae) of the Vistula excerpment in Warsaw. – Fragmenta PISARSKI, B. 1982: Ants (Hymenoptera, Formicoidea) of War- Faunistica 33: 109-128. saw and Mazovia. – Memorabilia Zoologica 36: 73-90. CZECHOWSKI, W., PISARSKI, B. & SKIBINSKA, E. 1979: Impact of PISARSKI, B. & CZECHOWSKI, W. 1978: Influence de la pression the urban pressure on the associations of social . – In- urbaine sur la myrmécofaune. – Memorabilia Zoologica 29: ternationales Symposium über Entomofaunistik in Mitteleu- 109-128. ropa, VII, 1977, Leningrad: 68-71 (in Russian). PISARSKI, B. & CZECHOWSKI, W. 1987: Structure and origin of ant CZECHOWSKI, W., RADCHENKO, A. & CZECHOWSKA, W. 2002: The communities of Warsaw. In: EDER, J. & REMBOLD, H. (Eds.): ants (Hymenoptera, Formicidae) of Poland. – Museum I In- Chemistry and biology of social insects. – Verlag J. Peperny, stytut Zoologii PAN, Warszawa, 200 + 1 pp. München, p. 605. DAUBER, J. 1995: Die Ameisen (Hymenoptera: Formicidae) von PISARSKI, B. & KUĹESZA, M. 1982: Characteristics of col- Mainz. – Unpublished Master thesis, Gutenberg-Universität onizing urban habitats. – Memorabilia Zoologica 37: 71-77. Mainz, Mainz, 150 pp. PISARSKI, B. & VEPSÄLÄINEN, K. 1987: Geographic variability of DAUBER, J. 1997: Ameisenfauna einer urbanen Landschaft. – Na- myrmecofauna of urban areas. In: EDER, J. & REMBOLD, H. turschutz und Landschaftsplanung 29: 303-309. (Eds.): Chemistry and biology of social insects. – Verlag J. Peperny, München, p. 604. DAUBER, J. & EISENBEIS, G. 1997: Untersuchungen zur Ameisen- fauna einer urbanen Landschaft am Beispiel der Stadt Mainz. READ, J.L. & ANDERSEN, A.N. 2000: The value of ants as early – Abhandlungen und Berichte des Naturkundemuseums Gör- warning bioindicators: responses to pulsed cattle grazing at an litz 69: 237-244. Australian arid zone locality. – Journal of Arid Environments 45: 231-251. GASPAR, CH. & THIRION, C. 1978: Modification des populations d'hyménoptères sociaux dans des milieux anthropogènes. – RESTREPO, C., ESPADALER, X. & DE HARO, A. 1985: Contribu- Memorabilia Zoologica 29: 61-77. ción al conocimiento faunístico de los formícidos de Garraf (Barcelona). – Orsis 1: 113-129. HOFFMANN, B.D. & ANDERSEN, A.N. 2003: Responses of ants to disturbance in Australia, with particular reference to functional RICHTER, K., KLAUSNITZER, B. & ZIMDARS, A. 1986: Zur Amei- groups. – Austral Ecology 28: 444-464. senfauna unterschiedlich urban beeinflußter Ruderalstellen im Gebiet von Leipzig (Hymenoptera, Formicidae). – Entomolo- HOFFMANN, B.D., GRIFFITHS, A.D. & ANDERSEN, A.N. 2000: Re- gische Nachrichten und Berichte 30: 115-120. sponses of ant communities to dry sulfur deposition from mining emissions in semi-arid tropical Australia, with implications SCHLICK-STEINER, B.C. & STEINER, F.M. 1999: Faunistisch-öko- for the use of functional groups. – Austral Ecology 25: 653-663. logische Untersuchungen an den freilebenden Ameisen (Hy- menoptera: Formicidae) Wiens. – Myrmecologische Nachrich- KLAUSNITZER, B. 1990: Ecology of urban fauna. – Mir, Moscow, ten 3: 9-53. 248 pp. (in Russian). SEIFERT, B. 1996: Ameisen: beobachten, bestimmen. – Naturbuch KONDOH, M. 1978: A comparison among ant communities in the Verlag, Augsburg, 352 pp. anthropogenic environment. – Memorabilia Zoologica 29: 79-92. STATSOFT, INC. 1995: STATISTICA for Windows [Computer KONDOH, M. & KITAZAWA, Y. 1984: Ant communities on the program manual]. Campus of UOEH and in an adjacent natural forest. – Journal of University of Occupational and Environmental Health 6: VANDERWOUDE, C., ANDERSEN, A.N. & HOUSE, A.P.N. 1997а: Ant 221-234. communities as Bio-indicators in relation to fire management of Spotted Gum (Eucaliptus maculate HOOK.) forests in South- KOVACHEV, А. 2005: The green system of Sofia. Urban aspects. – East Queensland. – Memoirs of the Museum of Victoria 56: PENSOFT Publishers, Sofia, Moscow, 368 pp (in Bulgarian). 671-675. LIPPKE, S. & CÖLLN, K. 1991: Über die Ameisen (Formicidae) VANDERWOUDE, C., ANDERSEN, A.N. & HOUSE, A.P.N. 1997b: von Köln. – Verhandlungen Westdeutscher Entomologentag Community organisation, biogeography and seasonality of ants 1990: 87-90. in an open forest of South-eastern Queensland. – Australian MCINTYRE, N.E. 2000: Ecology of urban : a review and Journal of Zoology 45: 523-537. a call to action. – Ecology and population biology 93: 825-835. VEPSÄLÄINEN, K. & PISARSKI, B. 1982: The structure of urban NEW, T.R. 2000: How useful are ant assemblages for monitor- ant communities along a geographical gradient from North Fin- ing habitat disturbance on grasslands in southeastern Austra- land to Poland. – Ossolineum ( in urban environment. lia? – Journal of Conservation 4: 153-159. Symposium in Warszawa Labłonna): 155-168. NUHN, T.P. & WRIGHT, C.G. 1979: An ecological survey of ants VEPSÄLÄINEN, K. & WUORENRINNE, H. 1978: Ecological effects (Hymenoptera: Formicidae) in a landscaped suburban habitat. of urbanization on the mound-building Formica L. species. – – American Midland Naturalist 102: 353-362. Memorabilia Zoologica 29: 191-202. PECK, S. L., MCQUAID, B. & CAMPBELL, C.L. 1998: Using ant YAMAGUCHI, T. 2004: Influence of urbanization on ant distribu- species (Hymenoptera, Formicidae) as a biological indicator tion in parks of Tokyo and Chiba City, Japan. – Ecological Re- of agroecosystem condition. – Environmental Entomology 27: search 19: 209-216. 1102-1110.

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